Nitrification in soils of secondary vegetational successions from Eucalyptus forest and grassland to cool temperate rainforest in Tasmania

Rates of nitrification in well drained granitic soils from forest stands and grassland of differing successional status and from beneath isolated individuals of several tree species were compared in a series of laboratory experiments. Fresh samples were perfused with distilled water or nutrient solution for 10 to 14 weeks at 20°C. The following treatments were applied to the soils singly and in combination: 200 and 400 g N g^–1 as (NH₄)₂SO₄; 100 g P g^–1 as KH₂PO₄; 4000 g CaCO₃ g^–1; inoculation of non-nitrifying soil with nitrifying soil; perfusion of nitrifying soil with leachate from non-nitrifying soil.Nitrification was absent or occurred at only a low rate in many soils; it generally increased as succession proceeded from nature grassland or eucalypt forest towards climax temperate rainforest, but decreased in mature climax forests. However, the influence of individual tree species was often paramount. Nitrification was stimulated by disturbance of a stand by disease. A possible inhibitor of nitrification in a rainforest soil could not be removed by leaching with water, nor transferred via the leachate to a nitrifying soil. Addition of P was without effect on either total amount of nitrate produced or on net mineralisation of soil N, but sometimes increased the rate of nitrification of added ammonium. Non-nitrifying rainforest soil of pH 4.3 was induced to nitrify only after addition of (NH₄)₂SO₄, inoculation with a nitrifying soil, and addition of CaCO₃ to raise pH by 3 units. However, once nitrification had commenced it could continue with little change in rate while pH decreased to a value of 3.4.It was concluded that rate of nitrification is dependent upon the presence of particular tree species in a stand, upon its history of disturbance, and hence in part upon the stand's successional status. It is not limited by pHper se within the range found in these soils, although an increase in pH may be necessary to initiate nitrification. In some soils the rate of nitrification may be limited by the level of ammonium substrate, and nitrifiers are virtually absent from others. Overall microbial activity is limited by lack of utilisable carbon substrate.     

A comparison of soil quality in adjacent cultivated,forest and native grassland soils

Changes in soil quality after 45 years of continuous production of corn (Zea mays L.) by the conventional tillage method (C) compared with adjacent poplar forest (F) and native grassland (G) sites were examined. The investigated parameters were: total and humified organic C, total N, light fraction content and composition, water-soluble organic C (WSOC), water-soluble carbohydrates (WSC), phenolic substances, biomass C, cumulative CO₂-C (soil respiration) (C _m), enzyme activities (alkaline phosphatase, protease, -glucosidase, urease, catalase and dehydrogenase). Empirical indexes of soil quality were also calculated: biomass C/organic C, specific respiration of biomass C (qCO₂), death rate quotient (qD), metabolic potential (MP), biological index of fertility (BIF), enzyme activity number (EAN) and hydrolysing coefficient (HC). Results indicate that long-term corn production at an intensive level caused a marked decline in all examined parameters. Between the undisturbed systems, native grassland showed higher values of soil quality parameters than forest site. The indexes most responsive to management practices that may provide indications of the effects of soil cultivation, as well as of the differently undisturbed ecosystems were: organic C, WSC, C _m, protease, -glucosidase, urease and HC. Soil enzyme activities were well related with, and not more sensitive than organic carbon.     

Effects of forest soil desiccation on the growth of Eucalyptus regnans F. Muell. seedlings

Abstract. The growth rate of Eucalyptus regnans seedlings in their first year can be much increased if the soil is first dried and then rewetted. The ratio of growth on predried soil to growth on undried soil (the Growth Ratio or GR) reaches a maximum at air-dryness (pF 6.0–6.4). In E. regnans forest soil, GR is greatest in humus-rich topsoil and declines with depth. The effect of air-drying persists for several months after rewetting when soil is stored under glasshouse conditions. It is largely unaffected by repeated drying and wetting, by the rate of drying or by the season of collection. The mixing of dried and undried soil or the placement of a layer of dried soil above undried soil produces an enhancement of growth proportional to the amount of dried soil added. Firing of a litter layer above soil at wilting point increases subsequent seedling growth to that in air-dried soil. The addition of ash from a litter fire to undried soil produces an increase in growth approximately equal to that caused by air-drying The drying effect is most pronounced in soils from mature E. regnans forest and nearby brackenland and is less in dense younger forest, frost-hollow grasslands and old grassy gaps in the mature forest. The effect is restored by the inoculation of E. regnans mycorrhizal roots from both dried and undried soil. The effect varies along an gradient from 500 to 1500 m a.s.l. and is a maximum in the wet E. regnans climatic zone and a minimum in zones or local aspects where forests are normally subject to frequent drying. The stimulatory effect on seedling growth in soils of the E. regnans zone may have an effect on the outcome of competition during regeneration in large gaps. Part of the growth responses previously ascribed to the ‘ash-bed’ effect may be due to the desiccation effect in these soils.     

Physicochemical and biological factors affecting atmospheric methane oxidation in gray forest soils

The decline of methane oxidizing activities in gray forest soil upon its conversion into arable land was shown to be caused by major changes in biotic and physicochemical properties of soil. Using the method of immune serums, methane-oxidizing bacteria were detected in both forest and agricultural soils, but their populations differed significantly in both abundance and composition. In the forest soil, the number of methanotrophs was an order of magnitude higher than in arable soil, amounting to 3.5 × 10⁸ and 0.24 × 10⁸ cells/g soil, respectively. All methane-oxidizing bacteria identified in the forest soil belonged to the genus Methylocystis, and 94% of these were represented by a single species, M. parvus. The arable soil was dominated by type I methanotrophs (Methylobacter and Methylomonas, 67.6%), occurring along with bacteria of the genus Methylocystis. In addition, arable soil is characterized by a low content of microbial biomass, lower porosity and water resistance of soil aggregates, and the predominance of nitrogen mineralization processes over those of nitrogen immobilization. These factors can also contribute to lower rates of methane oxidation in arable soil as compared to forest soil.__________Translated from Mikrobiologiya, Vol. 74, No. 2, 2005, pp. 255–260.Original Russian Text Copyright © 2005 by Kravchenko, Semenov, Kuznetsova, Bykova, Dulov, Pardini, Gispert, Boeckx, Cleemput, Galchenko.     

Carbon isotopic fractionation in subtropical brazilian grassland soils. Comparison with tropical forest soils

The natural relationship¹³C/¹²C determined in three soil profiles under grass vegetation indicated a depletion in organic¹³C at depth: theδ ¹³C was between −18‰ and −15‰ in the A horizons and ranged from −18 to −22‰ at depth. Previous work showed that in forest soils, whereδ ¹³C was near −28‰ in the upper horizon, there was, on the contrary, a relative enrichment of the lower strata. This meant thatδ ¹³C, initially different in the various topsoils, became more equal at depth. Comparison between dark, deep horizons (sombric horizons), which are certainly of illuvial origine, would confirm this:δ ¹³C of grassland and a forest sombric horizon were almost equal at around −22‰. These results might mean that, in natural ecosystems, the isotopic carbon composition of the soil underlying humus would be independent of the vegetation type. This would have practical implications for the use of¹³C as a tracer for soil organic matter studies.     

Root growth of subalpine and montane Eucalyptus seedlings at low soil temperatures

 This study examines the effect of different soil temperatures on root growth in seedlings of Eucalyptus pauciflora Sieber ex Sprengel subsp. pauciflora and Eucalyptus nitens (Deane & Maiden) Maiden. Seedlings were grown in a glasshouse in pots containing soil. Pots were held in water baths maintained at 3, 7 or 13°C, whilst shoots were exposed to ambient glasshouse temperatures. The experiments were designed to separate direct effects of soil temperature from effects due to differences in seedling size. In the first experiment, seedlings were grown to constant height (25 cm for both species), in the second to constant time (100 days for E. pauciflora and 64 days for E. nitens) and in the third experiment seedlings were transferred between soil temperatures. The rate of growth of both species increased with increasing soil temperature. E. nitens grew faster than E. pauciflora at 7 and 13°C, but E. pauciflora grew faster than E. nitens at 3°C. The rate of browning of roots increased with decreasing soil temperature and at a faster rate in E. nitens than E. pauciflora. Root length was highly correlated to root mass within diameter and colour classes (r² > 0.7). However, brown roots were heavier than white roots. Consequently, changes in root mass did not reflect changes in root length when the proportion of brown to white root also changed. For example, at a constant height of 25 cm at 3°C, E. nitens had greater root mass but lesser root length than E. pauciflora. E. pauciflora at 3°C grew faster, and had more root length and less brown roots than E. nitens. This supports the argument that E. pauciflora is better adapted than E. nitens to survive and grow at lower soil temperatures. Received: 16 December 1996 / Accepted: 2 April 1997     

Mineralogy of the rhizosphere in forest soils of the eastern United States

Chemical and mineralogical studies of forest soils from six sites in the northeastern and southeastern United States indicate that soil in the immediate vicinity of roots and fine root masses may show marked differences in physical characteristics, mineralogy and weathering compared to the bulk of the forest soil. Examination of rhizosphere and rhizoplane soils revealed that mineral grains within these zones are affected mechanically, chemically and mineralogically by the invading root bodies. In SEM/EDS analyses, phyllosilicate grains adjacent to roots commonly aligned with their long axis tangential to the root surface. Numerous mineral grains were also observed for which the edge abutting a root surface was significantly more fractured than the rest of the grain. Both the alignment and fracturing of mineral grains by growing roots may influence pedogenic processes within the rhizosphere by exposing more mineral surface to weathering in the root-zone microenvironment. Chemical interactions between roots and rhizosphere minerals included precipitation of amorphous aluminium oxides, opaline and amorphous silica, and calcium oxalate within the cells of mature roots and possible preferential dissolution of mineral grains adjacent to root bodies. Mineralogical analyses using X-ray diffraction (XRD) techniques indicated that kaolin minerals in some rhizosphere samples had a higher thermal stability than kaolin in the surrounding bulk forest soil. In addition, XRD analyses of clay minerals from one of the southeastern sites showed abundant muscovite in rhizoplane soil adhering to root surfaces whereas both muscovite and degraded mica were present in the immediately surrounding rhizosphere soil. This difference in mineral assemblages may be due to either K-enrichment in rhizoplane soil solutions or the preferential dissolution of biotite at the root-soil interface     

Contemporary carbon stocks of mineral forest soils in the Swiss Alps

Soil organic carbon (SOC) has been identified as the main globalterrestrial carbon reservoir, but considerable uncertainty remains as toregional SOC variability and the distribution of C between vegetationand soil. We used gridded forest soil data (8–km × 8–km)representative of Swiss forests in terms of climate and forest typedistribution to analyse spatial patterns of mineral SOC stocks alonggradients in the European Alps for the year 1993. At stand level, meanSOC stocks of 98 t C ha^–1 (N = 168,coefficient of variation: 70%) were obtained for the entiremineral soil profile, 76 t C ha^–1 (N =137, CV: 50%) in 0–30 cm topsoil, and 62 t Cha^–1 (N = 156, CV: 46%) in0–20 cm topsoil. Extrapolating to national scale, we calculatedcontemporary SOC stocks of 110 Tg C (entire mineral soil, standarderror: 6 Tg C), 87 Tg C (0–30 cm topsoil, standarderror: 3.5 Tg C) and 70 Tg C (0–20 cm topsoil, standarderror: 2.5 Tg C) for mineral soils of accessible Swiss forests(1.1399 Mha). According to our estimate, the 0–20 cm layers ofmineral forest soils in Switzerland store about half of the Csequestered by forest trees (136 Tg C) and more than five times morethan organic horizons (13.2 Tg C).At stand level, regression analyses on the entire data set yielded nostrong climatic or topographic signature for forest SOC stocks in top(0–20 cm) and entire mineral soils across the Alps, despite thewide range of values of site parameters. Similarly, geostatisticalanalyses revealed no clear spatial trends for SOC in Switzerland at thescale of sampling. Using subsets, biotic, abiotic controls andcategorial variables (forest type, region) explained nearly 60%of the SOC variability in topsoil mineral layers (0–20 cm) forbroadleaf stands (N = 56), but only little of thevariability in needleleaf stands (N = 91,R ² = 0.23 for topsoil layers).Considerable uncertainties remain in assessments of SOC stocks, due tounquantified errors in soil density and rock fraction, lack of data onwithin-site SOC variability and missing or poorly quantifiedenvironmental control parameters. Considering further spatial SOCvariability, replicate pointwise soil sampling at 8–km × 8–kmresolution without organic horizons will thus hardly allow to detectchanges in SOC stocks in strongly heterogeneous mountain landscapes.     

A comparative account of the microbiological characteristics of soils under natural forest,grassland and cropfield from Eastern India

Microbiological and physico-chemical characteristics of tropical forest, grassland and cropfield soils from India were investigated. The study revealed that the conversion of natural forest led to a reduction of soil organic C (26–36%), total N (26–35%), total P (33–44%), microfungal biomass (44–66%) and total microbial biomass C, N and P (25–60%) over a period of 30–50 years. Comparative analysis of microbial activity in terms of basal soil respiration revealed maximum activity in the forest and minimum in the cropfield soil. Analysis of microbial metabolic respiratory activity (qCO₂) indicated relatively greater respiratory loss of CO₂-C per unit microbial biomass in cropfield and grassland than in forest soil. Considering the importance of the microbial component in soil, we conclude that the conversion of the tropical forest to different land uses leads to the loss of biological stability of the soil.     

A comparison of arbuscular and ectomycorrhizal Eucalyptus coccifera: growth response, phosphorus uptake efficiency and external hyphal production

Eucalyptus coccifera Hook., a plant capable of forming both arbuscular mycorrhizas and ectomycorrhizas, was used to compare the effects of the two mycorrhizal types on phosphorus uptake and C allocation. Seedlings were grown in a P-deficient soil/sand mixture inoculated with peat/vermiculite spawn of Laccaria bicolor (Maire) Orton or Thelephora terrestris (Ehrh.) Fr.; or with 250-μm sievings from leek colonized by Glomus caledonium (Nicol. & Gerd.) Trappe & Gerde., Glomus sp. type E3 or Glomus mosseae (Nicol. & Gerd.) Gerd. & Trappe or with autoclaved spawn (non-mycorrhizal control). Before the 89-d harvest, a subset of the harvested plants was labelled with ¹⁴C (45–60-min pulse, 202-h chase). Growth promotion and the increase in seedling P content was largest in the two ectomycorrhizal treatments. Production of fluorescein diacetate-stained external hyphae was three to seven times higher by ectomycorrhizal (ECM) fungi compared with arbuscular mycorrhizal (AM) fungi and was highly correlated with P uptake and shoot weight. Phosphorus inflow rates of ECM and AM seedlings were 3·8 times, and 2·0–2·7 times those of non-mycorrhizal seedlings. Phosphorus acquisition efficiencies were similar (11·2 and 10·0 μmol P mmol⁻¹C for T. terrestris and Glomus E3 plants, respectively) for the two mycorrhizal types, and appeared to be greater than in uninoculated plants (7.2 μmol P mmol⁻¹C) grown at the same P level.     

Diagnosis of zinc deficiency in seedlings of a tropical eucalypt (Eucalyptus urophylla S. T. Blake)

A glasshouse experiment was undertaken to establish the internal Zn requirement for shoot growth of Eucalyptus urophylla, a fast-growing commercial plantation species widely planted in tropical regions of the world. A Zn-deficient sand was supplemented with ten rates of Zn and seedlings were harvested after three months. In Zn-deficient plants the new growth was dwarfed with small, necrotic leaves and short internodes. Foliar Zn concentrations declined markedly with leaf age in both Zn-deficient and Zn-adequate plants. The critical Zn concentration for the diagnosis of Zn deficiency also fell with leaf age. Zinc concentrations in the youngest fully expanded leaf ranged from 8–11 g Zn g^–1 dry weight in plants with severe symptoms to 30–37 g Zn g^–1 dry weight in non-deficient plants. The critical Zn concentration for the diagnosis of Zn deficiency at 90% of maximum shoot growth in the same leaf was 21 g Zn g^–1 dry weight. This value is nearly twice that reported for several other species of eucalypts and may indicate a higher internal demand for Zn in tropical than in temperate eucalypts.     

Denitrification in the top and sub soil of grassland on peat soils

Denitrification is an important process in the nitrogen (N) balance of intensively managed grassland, especially on poorly drained peat soils. Aim of this study was to quantify the N loss through denitrification in the top and sub soil of grassland on peat soils. Sampling took place at 2 sites with both control (0 N) and N fertilised (+ N) treatments. Main difference between the sites was the ground water level. Denitrification was measured on a weekly basis for 2 years with a soil core incubation technique using acetylene (C₂H₂) inhibition. Soil cores were taken from the top soil (0–20 cm depth) and the sub soil (20–40 cm depth) and incubated in containers for 24 hours. The denitrification rate was calculated from the nitrous oxide production between 4 and 24 hours of incubation. Denitrification capacities of the soils and the soil layers were also determined.The top soil was the major layer for denitrification with losses ranging from 9 to 26 kg N ha^–1 yr^–1 from the O N treatment. Losses from the top soil of the + N treatment ranged from 13 to 49 kg N ha^–1 yr^–1. The sub soil contributed, on average, 20% of the total denitrification losses from the 0–40 layer. Losses from the 0–40 cm layer were 2 times higher on the + N treatment than on the O N treatment and totalled up to 70 kg N ha^–1 yr^–1. Significant correlation coefficients were found between denitrification activity on the one hand, and ground water level, water filled pore space and nitrate content on the other, in the top soil but not in the sub soil. The denitrification capacity experiment showed that the availability of easily decomposable organic carbon was an important limiting factor for the denitrification activity in the sub soil of these peat soils.     

Tree growth and soil acidification in response to 30 years of experimental nitrogen loading on boreal forest

Relations among nitrogen load, soil acidification and forest growth have been evaluated based on short‐term (<15 years) experiments, or on surveys across gradients of N deposition that may also include variations in edaphic conditions and other pollutants, which confound the interpretation of effects of N per se. We report effects on trees and soils in a uniquely long‐term (30 years) experiment with annual N loading on an un‐polluted boreal forest. Ammonium nitrate was added to replicated (N=3) 0.09 ha plots at two doses, N1 and N2, 34 and 68 kg N ha^?1 yr^?1, respectively. A third treatment, N3, 108 kg N ha^?1 yr^?1, was terminated after 20 years, allowing assessment of recovery during 10 years. Tree growth initially responded positively to all N treatments, but the longer term response was highly rate dependent with no gain in N3, a gain of 50 m³ ha^?1 stemwood in N2 and a gain of 100 m³ ha^?1 stemwood in excess of the control (N0) in N1. High N treatments caused losses of up to 70% of exchangeable base cations (Ca²⁺, Mg²⁺, K⁺) in the mineral soil, along with decreases in pH and increases in exchangeable Al³⁺. In contrast, the organic mor‐layer (forest floor) in the N‐treated plots had similar amounts per hectare of exchangeable base cations as in the N0 treatment. Magnesium was even higher in the mor of N‐treated plots, providing evidence of up‐lift by the trees from the mineral soil. Tree growth did not correlate with the soil Ca/Al ratio (a suggested predictor of effects of soil acidity on tree growth). A boron deficiency occurred on N‐treated plots, but was corrected at an early stage. Extractable NH₄⁺ and NO₃^?were high in mor and mineral soils of on‐going N treatments, while NH₄+ was elevated in the mor only in N3 plots. Ten years after termination of N addition in the N3 treatment, the pH had increased significantly in the mineral soil; there were also tendencies of higher soil base status and concentrations of base cations in the foliage. Our data suggest the recovery of soil chemical properties, notably pH, may be quicker after removal of the N‐load than predicted. Our long‐term experiment demonstrated the fundamental importance of the rate of N application relative to the total amount of N applied, in particular with regard to tree growth and C sequestration. Hence, experiments adding high doses of N over short periods do not mimic the long‐term effects of N deposition at lower rates.     

Factors affecting the initiation and growth of aboveground adventitious roots in a tropical cloud forest tree: an experimental approach

Some of the proximate factors that would induce aboveground stems to produce adventitious roots were investigated experimentally on Senecio cooperi, a tropical cloud forest tree. Stem segments were air-layered with different treatments to promote root formation, and the number of roots initiated and rates of root growth were monitored for 20 weeks. Treatments were the application of wet epiphytes or dry epiphytes plus associated humus, sponges wetted with either water or nutrient solutions, or dry sponges. Controls (stem segments with nothing applied) were also monitored. Numbers of adventitious roots formed and rates of subsequent root growth differed among treatments. Wet epiphyte/humus and nutrient solutions were most effective in producing roots, which suggests that epiphytes and the nutrients they intercept and retain within the canopy may cue adjacent host tissue to exploit this resource.     

The soil seed bank and understorey regeneration in Eucalyptus regnans forest,Victoria

Abstract The soil seed bank and its relation to the extant vegetation in a Eucalyptus regnans F. Muell. forest in the Central Highlands of Victoria were examined. The average seed density was 430 germinable seeds m^?2 to a depth of 2 cm. There was a polynomial regression relationship between the density and species richness of seeds in soil and forest age (0. 6–54 years). Species richness was not significantly different among soil depths (0- 2 , 2- 5 , 5–10 and 10–20 cm) in the forest stand of 54 years old. More seeds germinated from the 5–10 cm depth than from the other depths. Forbs accounted for 73% of the total germinable seeds and there was no germination of E. regnans. The number of species, particularly woody plant species, germinating from the soil seed bank were significantly lower than in the extant vegetation. However, almost all species present in the soil seed bank were present in the vegetation. The soil seed bank provides an important source for the rapid regeneration of understorey vegetation following clear-cutting and slash-burning in the E. regnans forest. The rapid understorey establishment may play an important role in protecting soil from erosion, in nutrient conservation, replacement and redistribution. The soil seed bank may also be a necessary source of maintaining genetic diversity in the forest over the long term.     

Quantification of ammonium sorption in acid forest soils by sorption isotherms

Quantity/Intensity relations of NH₄ ⁺ were established to evaluate the role of NH₄ ⁺ sorption as a N-sink as well as for the transport of NH₄ ⁺ within the soil profile. Two haplic podzols (originating from Phyllite and Granite) from the German Fichtelgebirge were used. Exchange isotherms were obtained from batch experiments with NH₄ ⁺ concentrations between 0.01 to 0.6 mmol NH₄ ⁺ L^-1. Background solutions had a composition close to the natural soil solutions to identify the cations displaced by NH₄ ⁺. Organic layers sorbed a maximum amount of 0.3–0.5 mmol_c NH₄ ⁺ 100 g^-1, A horizons 0.1–0.2 mmol_c NH₄ ⁺ 100 g^-1 and B horizons 0.09 mmol_c 100 g^-1, corresponding to less than 1% of the cation exchange capacities. NH₄ ⁺ displaced K⁺ and Ca²⁺ throughout the whole profile, and additionally Al in the subsoil horizons and H₃O⁺ in the organic and A horizons. Initial mass isotherms (Quantity/Quantity isotherms) were established with distribution coefficients ranging from 0.17 to 0.35 for NH₄ ⁺ with highest values in the O horizons. While exchangeable NH₄ ⁺ in acid forest soils is considered to be only a minor long-term sink for deposited NH₄ ⁺, it has its relevance in the seasonal dynamics and transport of NH₄ ⁺.     

The effect of regeneration burns on the growth,nutrient acquisition and mycorrhizae of Eucalyptus regnans F. Muell. (mountain ash) seedlings

This study was conducted to compare the effects on the growth of Eucalyptus regnans seedlings of unheated soil and soil heated to different extents (as indicated by soil colour–bright red or black) in burnt logging coupes, and to separate the effects of heating of the soil on direct nutrient availability and on morphotypes and effectiveness of ectomycorrhizae. Burnt soils were collected from three logging coupes burnt 2, 14 and 25 months previously and unbumt soil from adjacent regrowth forests. Compared to unburnt soil, the early seedling growth was stimulated in black burnt soil from all coupes (burnt 2, 14 and 25 months previously). Seedling growth was generally poor in red burnt soil, especially in soil collected 2 months after burning. However, the concentration of extractable P was extremely high in red burnt soil, especially in soil collected 2 months after burning. In black burnt soil, extractable P was increased in soil 2 months after burning, but not in the soils collected 14 or 25 months after burning. However, both total P content and concentration in seedlings were increased in all collections of black burnt soil. Frequency of ectomycorrhizae was high in seedlings grown in all black burnt soils, but the mycorrhizal mantles were poorly developed in seedlings in black burnt soil collected 2 months after burning. Seedlings were also ectomycorrhizal in red burnt soil, except in soil collected 2 months after burning. Fine root inocula from seedlings grown in black burnt soils collected 14 and 25 months after burning significantly stimulated both seedling growth and P uptake compared with the uninoculated control, whereas the fine root inocula from the seedlings grown in all the other soils did not. These results suggest that, in black burnt soil, both direct nutritional changes and changes in the ectomycorrhizae may contribute to seedling growth promotion after regeneration burns. The generally poor seedling growth in red burnt soils is likely to have been due to N deficiency as the seedlings in these soils were yellow-green and the tissue concentrations of N were significantly lower than in other treatments. This revised version was published online in June 2006 with corrections to the Cover Date.     

Comparison of ecosystem water-use efficiency among Douglas-fir forest, aspen forest and grassland using eddy covariance and carbon isotope techniques

Comparisons were made among Douglas‐fir forest, aspen (broad leaf deciduous) forest and wheatgrass (C₃) grassland for ecosystem‐level water‐use efficiency (WUE). WUE was defined as the ratio of photosynthetic CO₂ assimilation rate and evapotranspiration (ET) rate. The ET data measured by eddy covariance were screened so that they overwhelmingly represented transpiration. The three sites used in this comparison spanned a range of vegetation (plant functional) types and environmental conditions within western Canada. When compared in the relative order Douglas‐fir (located on Vancouver Island, BC), aspen (northern Saskatchewan), grassland (southern Alberta), the sites demonstrated a progressive decline in precipitation and a general increase in maximum air temperature and atmospheric saturation deficit (D_max) during the mid‐summer. The average (±SD) WUE at the grassland site was 2.6±0.7 mmol mol^?1, which was much lower than the average values observed for the two other sites (aspen: 5.4±2.3, Douglas‐fir: 8.1±2.4). The differences in WUE among sites were primarily because of variation in ET. The highest maximum ET rates were approximately 5, 3.2 and 2.7 mm day^?1 for the grassland, aspen and Douglas‐fir sites, respectively. There was a strong negative correlation between WUE and D_max for all sites. We also made seasonal measurements of the carbon isotope ratio of ecosystem respired CO₂ (δ_R) in order to test for the expected correlation between shifts in environmental conditions and changes to the ecosystem‐integrated ratio of leaf intercellular to ambient CO₂ concentration (c_i/c_a). There was a consistent increase in δ_R values in the grassland, aspen forest and Douglas‐fir forest associated with a seasonal reduction in soil moisture. Comparisons were made between WUE measured using eddy covariance with that calculated based on D and δ_R measurements. There was excellent agreement between WUE values calculated using the two techniques. Our δ_R measurements indicated that c_i/c_a values were quite similar among the Douglas‐fir, aspen and grassland sites, despite large variation in environmental conditions among sites. This implied that the shorter‐lived grass species had relatively high c_i/c_a values for the D of their habitat. By contrast, the longer‐lived Douglas‐fir trees were more conservative in water‐use with lower c_i/c_a values relative to their habitat D. This illustrates the interaction between biological and environmental characteristics influencing ecosystem‐level WUE. The strong correlation we observed between the two independent measurements of WUE, indicates that the stable isotope composition of respired CO₂ is a useful ecosystem‐scale tool to help study constraints to photosynthesis and acclimation of ecosystems to environmental stress.